title: A pollinator shift explains floral divergence in an orchid species complex in South Africa.
abstract: OBJECTIVE
  Floral diversification driven by shifts between pollinators has been one of the key explanations for the radiation of angiosperms. According to the Grant-Stebbins model of pollinator-driven speciation, these shifts result in morphologically distinct 'ecotypes' which may eventually become recognizable as species. The current circumscription of the food-deceptive southern African orchid Eulophia parviflora encompasses a highly variable monophyletic species complex. In this study, two forms were identified within this complex that differ in distribution, floral morphology, scent chemistry and phenology, and a test was made of whether these differences represent adaptations for different pollinators.
  
  
  RESULTS
  Multivariate analysis of floral and vegetative traits revealed that there are at least two discrete morphological forms in the species complex. Field observations revealed that each form is pollinated by a different insect species, and thus represent distinct ecotypes. The early-flowering coastal form which has long spurs and floral scent dominated by sesquiterpene compounds is pollinated exclusively by the long-tongued bee Amegilla fallax (Apidae, Anthophorinae), while the late-flowering inland form with short spurs and floral scent dominated by benzenoid compounds is pollinated exclusively by the beetle Cyrtothyrea marginalis (Cetoniinae; Scarabaeidae). Choice experiments in a Y-maze olfactometer showed that beetles are preferentially attracted to the scent of the short-spurred form. A spur-shortening experiment showed that long spurs are required for effective pollination of the bee-pollinated form. Although it was initially thought likely that divergence occurred across a geographical pollinator gradient, plants of the long-spurred form were effectively pollinated when transplanted to an inland locality outside the natural coastal range of this form. Thus, the underlying geographical basis for the evolution of ecotypes in the E. parviflora complex remains uncertain, although early flowering in the long-spurred form to exploit the emergence of naïve bees may restrict this form to coastal areas where there is no frost that would damage flower buds. Later flowering of the short-spurred form coincides closely with the emergence of the pollinating beetles following winter frosts.
  
  
  CONCLUSIONS
  This study identifies a shift between bee and beetle pollination as the main driver of floral divergence in an orchid species complex. Floral scent and spur length appear to be key traits in mediating this evolutionary transition.
authors.affiliation: School of Life Sciences, University of KwaZulu-Natal, Private Bag X01, Scottsville, 3209, South Africa.
authors.forename: Craig I
authors.initials: CI
authors.lastname: Peter
authors.forename: Steven D
authors.initials: SD
authors.lastname: Johnson
mesh_terms: D000818:Animals; D001516:Bees / Q000502:physiology*; D005075:Biological Evolution; D001947:Breeding; D002755:Choice Behavior; D001517:Coleoptera / Q000502:physiology*; D005426:Flight, Animal; D035264:Flowers / Q000033:anatomy & histology / Q000502:physiology*; D005843:Geography; D009812:Odorants; D064367:Olfactometry; D029595:Orchidaceae / Q000033:anatomy & histology / Q000502:physiology*; D010858:Pigmentation; D011058:Pollen / Q000502:physiology; D025341:Principal Component Analysis; D013019:South Africa; D013997:Time Factors
keywords: Amegilla; Cetoniinae; Eulophia; Grant–Stebbins model; Orchidaceae; Y-maze olfactometer; bee pollination; beetle pollination; phenology; pollination ecotypes; pollinator-driven speciation; scent